J Mammal Evol

DOI 10.1007/s10914-015-9307-8

ORIGINAL PAPER

A New Species of Prolibytherium (Ruminantia, Mammalia)

from Pakistan, and the Functional Implications of an Atypical
Atlanto-Occipital Morphology
Melinda Danowitz 1 & Rebecca Domalski 1 & Nikos Solounias 1

# Springer Science+Business Media New York 2015

Abstract We describe a new species of Prolibytherium, P.

fusus, sp. nov., from the lower Miocene of Pakistan, thus extending the genus to Asia. Prolibytherium is otherwise known
only from Libya. This species differs from Prolibytherium
magnieri in several basioccipital and atlanto-occipital morphologies. Namely, the posterior basioccipital tuberosities are continuous at the midline and lack the elevated transverse ridge
seen in P. magnieri, and the notch formed between the lateral
occipital condyles and paraoccipital process is lower. Both species of Prolibytherium have a characteristic ventrally fused occipital condyle at the midline, with a notably fuller circumferential articular surface. Prolibytherium magnieri also has thickened dorsal and ventral arches of the atlas. These specimens
also possess a longitudinal groove for the Eustachian tube extending from the alisphenoid canal to the bullae, and a second
deep grove isolating the basisphenoid bone from the temporal
bone. These, plus several other atlanto-occipital morphologies
strengthen the cervical support of the head. This is especially
important for Prolibytherium, as the taxon possesses massive
aliform cranial appendages. We relate the approximation of the
occipital condyles to a convergent state in two giraffids
(Giraffokeryx punjabiensis and Schansitherium tafeli), each of
which possesses multiple pairs of ossicones, presumably necessitating a strengthened atlanto-occipital joint.

Keywords Prolibytherium . Atlanto-occipital joint .

Basicranium . Ruminantia . Cranial appendages

* Melinda Danowitz
mdanowit@nyit.edu
1

Department of Anatomy, New York Institute of Technology College

of Osteopathic Medicine, Old Westbury, NY 11568, USA

Introduction
Prolibytherium magnieri is a specialized Miocene ruminant
with massive flattened cranial appendages, and a specialized
atlanto-occipital joint. Arambourg described it from Gebel
Zelten (16 Ma) in North Africa in 1961, and Hamilton
reviewed and described it in more detail in 1973. It is known
only from the lower Miocene of Gebel Zelten of Libya.
Prolibytherium magnieri is the type species for
Prolibytherium, and is presently the only species described
for the genus. We introduce a new species of Prolibytherium
found in the Zinda Pir of Pakistan, the first introduction of the
genus in Asia. The establishment of a new and geologically
older (19 Ma) species is based on differences in basicranial
morphology from P. magnieri.
The phylogeny of these ruminants is currently unresolved;
Prolibytherium has been placed in Sivatheriidae (Hamilton
1973), Palaeomerycidae (Janis and Scott 1987; Prothero and
Liter 2007; Solounias 2007), and Climacoceratidae (Pickford
et al. 2001; Morales et al. 2003; Snchez et al. 2010). We
briefly review the systematic affiliation of Prolibytherium,
and provide arguments strengthening its association with
Climacoceratidae.
The unique ventral fusion of the occipital condyles in
Prolibytherium has been mentioned briefly by Snchez et al.
(2010) and Solounias (2007), but the implications on the support and range of motion of the neck have yet to be discussed.
The cranial appendages of Prolibytherium are notably atypical
for ruminants, as they form a massive, slightly concave plate,
which is oriented parallel to the dorsal aspect of the skull
(Snchez et al. 2010). Solounias (2007) suggested that these
cranial appendages encompassed two pairs of structures (anterior and posterior), which merge and form a common web at
the base. Snchez et al. (2010) described the appendages as
having an asymmetric Bx^ shaped scaffold, which supports

J Mammal Evol

the lateral bony expansions. Hamilton (1973) described these

as large and aliform, and fused to the frontal and parietal bones
with no visible suture at the base. We relate these uniquely
structured and immense appendages to the atypical morphology of the atlanto-occipital joint.

Materials and Methods

50 mm

Prolibytherium magnieri
Fairly complete specimen (braincase with cranial appendages): NHM UK PVM21901
Fragmentary specimens, with fairly complete braincases:
NHM UK PVM99897, NHM UK PVM99896a, NHM UK
PVM26679
Atlas: NHM UK PVM99896b

Prolibytherium fusus, sp. nov.

Braincase with cranial appendages broken off: PMNH Z 162
The original specimens were examined for morphological
characteristics and anatomical descriptions.
Institutional abbreviationsAMNH, American Museum
of Natural History, New York, New York, USA; PMNH, The
Pakistan Natural History Museum, Islamabad, Pakistan; NHM
UK, The Natural History Museum, London, U.K; HPM,
Hezheng Paleozoological Museum, Hezheng, China.

30 mm

Fig. 1 Prolibytherium magnieri a, massive aliform cranial appendages

(NHM UK PVM21901) and their b, internal structure (NHM UK
PVM99896a)

Systematic Paleontology
20 mm

Order Artiodactyla Owen, 1848

Family cf. Climacoceratidae Hamilton, 1978
Genus Prolibytherium Arambourg, 1961

1
2
3

Type species Prolibytherium magnieri Arambourg, 1961

Generic DiagnosisMedium-sized (similar in size to
Rangifer tarandus) with flat, aliform cranial appendages oriented horizontal to the braincase, with the anterior and posterior pairs merged at a webbed base (Fig. 1a). The anterior pair
is smaller than the posterior pair. The cranial appendages grow
from the frontal bone, and are situated posterior to the supraorbital foramina. Their surface is smooth with shallow, thin
vascular impressions. Internally, the cranial appendages appear to have a medulla of cancellous bone surrounded by a
thick cortex (Fig. 1b). The maximum thickness of the cranial
appendages is ~300 mm. There are two lacrimal foramina at
the edge of the orbit. Ventrally, the occipital condyles are
fused at the median plane (Fig. 2). The posterior basioccipital
tuberosities are separated from the condyles by a deep, narrow
groove. The Eustachian tube left a long impression along the

5
Fig. 2 Prolibytherium magnieri braincase (NHM UK PVM99897) in
ventral view. (1)the longitudinal groove extending from the level of
the alisphenoid to the bullae for the Eustachian tube. (2)the medial deep
groove extending from the level of the alisphenoid to the posterior
basioccipital tuberosities that separates the basioccipital from the
temporal bone. (3)the posteriorly-directed posterior basioccipital tuberosities. (4)the ventrally fused occipital condyles characteristic of
Prolibytherium. (5)the narrow groove separating the approximated
basioccipital tuberosities

J Mammal Evol

basisphenoid bone, extending from the level of the alisphenoid

canal to the bullae. In addition, there is a second, more medial
groove between the Eustachian groove and the lateral edge of
the basioccipital-basisphenoid. This groove on the
basisphenoid extends from the alisphenoid to the posterior
basioccipital tuberosities, and separates the basisphenoid from
the temporal bone. The bullae are large and spherical. The
dorsal and ventral arches of the atlas are thickened (Fig. 3).
The premolars and molars are brachydont, and the upper P2
is slightly longer than the P3. The masseteric insertion on the
mandible forms a distinct notch from the body of the mandible.
This notch is anterior to the fossa for the masseter profundus,
and the greatest depth of the notch is in the same plane as the
superior margin of the foramen magnum. The mastoid process
is situated in the same plane as the occipital. Hamilton (1973)
included a more complete description of the genus.

Prolibytherium magnieri Arambourg, 1961

Type LocalityGebel Zelten, Libya (middle Miocene)
HolotypeA cranium with badly shattered cranial appendages described by Arambourg (1961).
Referred SpecimensHamilton (1973) listed 34 specimens from Natural History Museum-London, Musum
National dHistoire Naturelle- Paris, and Museum of
Paleontology University of California Berkley. This included
an almost complete skull, cranial fragments, several dentitions
and mandibles, and numerous postcranial specimens.
Diagnosis of type speciesAs for the genus.
CommentsTwo studies have focused on P. magnieri.
Hamilton (1973) described in detail the material, and
Snchez et al. (2010) reviewed Prolibytherium and proposed
that sexual dimorphism existed, with females possessing cranial appendages that retain the same pattern but are thin and
cylindrical and without the webbed aliform connection. Barry
et al. (2005) briefly compared Prolibytherium to Progiraffa,
and they referred a Zinda Pir braincase (PMNH Z 162), along
with a large sample of fragmentary specimens, to Progiraffa.
We are currently reassigning this braincase to Prolibytherium.
Atlas (Fig. 3) The dorsal and ventral arches are thickened.
The cranial articular facets, which articulate with the occipital condyles, are deep with a strongly bent lateral boundary. The facets are joined with a thickened bony connection
on the dorsal and ventral surfaces. There is a widened notch
formed on the lateral edge between the dorsal and ventral
aspects of the cranial articular facet. This notch articulates
with the junction between the posterior and ventral surfaces
of the occipital condyles. The ventral tubercle is strong,
and forms a large and thickened protrusion on the caudal aspect of the ventral arch. The ventral tubercle extends from the posterior edge of the vertebra and the
caudal articular facets as a triangular bony expansion.

50 mm
Fig. 3 Prolibytherium magnieri atlas (NHM UK PVM99896b) in a,
dorsal and b, caudal views

The anterior edge of the dorsal and ventral arches forms

a U-shape with a small notch at the midline on the
ventral surface. The dorsal arch has a median longitudinal keel, with a double bony protrusion on the posterior
end. The caudal articular facets are oval shaped, widest
medially, and are concave laterally. The caudal articular
facet extends laterally onto the postero-lateral process of
the alar wing.
Prolibytherium fusus, sp. nov.
(Fig. 4)
EtymologyFrom the Latin fusus Bthe fused one,^ in reference to the characteristic fused occipital condyles
HolotypePMNH Z 162, caudoventral portion of
braincase
Locality and Geological AgeLocality Z 124 in the
Vihowa Formation of Pakistan (early Miocene). Locality Z
124 is located at the top of a normal polarity zone that has
been correlated with the 6n polarity interval and would be
slightly older than 18.7 Ma (Lindsay et al. 2005).
Differential DiagnosisProlibytherium fusus differs from
P. magnieri in the following characters: the anterior
basioccipital tuberosities are less distinct, and their surface
contains longitudinal ridges instead of small bumps seen on
P. magnieri. The elongated fossa between the posterior and
anterior basioccipital tuberosities is one unified surface in

J Mammal Evol

20 mm

the paraoccipital process and the lateral occipital condyles is

thicker and lower in P. fusus (Fig. 4: character 5).
DescriptionThe Zinda Pir specimen PMNH Z 162 is a
partial braincase. The cranial appendages are broken off. A
more complete description is in Barry et al. (2005).

a
3

Description of the Prolibytherium magnieri

and P. fusus Basicranium and Occipital Condyles
Anterior Basioccipital Tuberosities These are slight ovoid
bony swellings, which are separated at the midline.

1
7
2

Fossa Between Anterior and Posterior Basioccipital Tuberosities, and Shape of Basioccipital Bone The fossa is
concave and contains longitudinal striations. There is a deep
longitudinal groove on the basisphenoid extending from the
alisphenoid to the posterior basioccipital tuberosities that separates the basioccipital from the temporal bone (Figs. 2 and 4:
character 2). Lateral to this, there is also a distinct groove on
the basisphenoid from the alisphenoid to the bullae for the
Eustachian tube (Figs. 2 and 4: character 1).

6
20 mm
Fig. 4 Prolibytherium fusus, sp. nov. braincase (PMNH Z 162) in a,
ventral, b, left posterolateral, and c, posterior views. (1)the
longitudinal groove extending from the level of the alisphenoid to the
bullae for the Eustachian tube. (2)the medial deep groove extending
from the level of the alisphenoid to the posterior basioccipital tuberosities
that separates the basioccipital from the temporal bone. (3)the
posteriorly-directed posterior basioccipital tuberosities. (4)the
ventrally fused occipital condyles characteristic of Prolibytherium.
(5)the narrow groove separating the approximated basioccipital
tuberosities. (6)the low-positioned notch between the paraoccipital
process and the occipital condyle. (7)the alisphenoid canal

P. magnieri, and is separated into two plates by a midline

longitudinal groove in P. fusus. In P. fusus, the posterior
basioccipital tuberosities are more approximated with the articular surface of the occipital condyles, and are thicker and
shorter. The posterior basioccipital tuberosities are also approximated across the midline in P. fusus, separated only by
a narrow, deep groove, and the caudal surface contains several
bony growths concentrated medially. In P. magnieri, the posterior basioccipital tuberosities are separated at the midline,
also by a deep groove, and the surface contains a distinct
elevated transverse ridge, and no bony growths. The Ushaped ventral margin on the occipital condyles is more shallow in P. fusus, and the condyles are oriented laterally, whereas the condyles are oriented dorso-laterally and have a deeper
U-shaped ventral margin in P. magnieri. The notch between

Posterior Basioccipital Tuberosities The posterior

basioccipital tuberosities are positioned rostral to the occipital
condyles, and are separated from the condyles by a distinct deep
groove. The articular surface of the condyles is not continuous
with the tuberosities. The posterior basioccipital tuberosities are
prominent, thick bony protrusions, which are approximated
near the midline. The tuberosities are angled slightly posteriorly
(Figs. 2 and 4: character 3), and follow the orientation of the
occipital condyles. On two of the four P. magnieri specimens,
the medial surface of the tuberosities expands rostrally, forming
a triangular extension onto the basioccipital.
Occipital Condyles (Ventral View) The two condyles are
thickened and fused at the midline, with no visible fusion line,
forming one unified articular surface. This forms a U-shaped
ventral margin to the foramen magnum. The surface of the
fused condyles is full, rounded, and bulging. The junction
between the posterior and ventral articular surfaces is rounded
and not distinct. There is a notch on the antero-dorsal aspect of
the lateral surface of the condyle. The fusion of the occipital
condyles makes these taxa unique among artiodactyls.
Occipital Aspect of the Skull The posterodorsal articular
surface of the occipital condyles is expanded. The notch
formed between the paroccipital process and condyle is
displaced ventrally, so that the notch is more ventral than the
dorsal edge of condyle. The occipital surface is on the same
plane as the mastoid bone and paroccipital process. The attachment for the nuchal ligament, the external occipital

J Mammal Evol

protuberance, has a central elevated ridge on the midline.

There are two deep pits positioned lateral to the ridge.

the Eustachian tube is elongated and horizontal, and embedded lateral to this groove. This Eustachian tube morphology
seen both in Prolibytherium and the Z 162 braincase is unlike
that of other ruminants.

Discussion
Systematic Position of Prolibytherium
Assignment of the Zinda Pir Braincase to Prolibytherium
Barry et al. (2005) referred the braincase PMNH Z 162 from
Zinda Pir to Progiraffa exigua, as well as numerous teeth, a
maxilla, postcranial elements and an isolated cranial
appendage. Pilgrim (1908) established Progiraffa exigua
based on a left second and third molar from the Upper Nari
beds (now the Chitarwata Formation [Antoine et al. 2013]) of
Dako Nala in Bugti Hills of Baluchistan (Pakistan). The specimen was figured in Pilgrim (1911: pl. I, fig. 1) together with a
second lower molar (pl. I, fig. 2) identified as Progiraffa sp.
from the Blower Siwaliks^ of Sind. A third lower molar (pl. I,
fig. 3) and two upper molars were identified as Progiraffa
sivalensis from the lower Siwaliks of the Potwar (Pilgrim
1908). These teeth are all primitive-looking giraffid teeth;
the protoconid, metaconid, and the other cuspids form walls
that are buccally and lingually inclined and slightly convex,
rather than straight and vertical.
The basis of the referral by Barry et al. (2005) was size,
geographic proximity, and the age of the fossils. While it is
common in the Miocene to find three or more giraffid species
per locality, and thus there may have been multiple species of
large ruminants in the lower Miocene of southern Asia, we
refer the Zinda Pir braincase to Prolibytherium. At this time,
we do not exclude the possibility of the presence of Progiraffa
in the Zinda Pir collection. We simply exclude the braincase
from the remainder of the Progiraffa sample, because it better
matches Prolibytherium. The new identification extends the
range of Prolibytherium into Asia and strengthens faunal similarities between southern Asia and northern Africa.
Barry et al. (2005) noted the distinct thickness of the dorsal
aspect of the Z 162 braincase, suggesting the presence of large
cranial appendages as are seen in Prolibytherium. The broken
edge of the cranial appendages is posterior and covers the
width of the calvaria confirming the presence of structures
similar to those of P. magnieri. In addition, we note three
additional characters that place the braincase closer to
Prolibytherium than Progiraffa. First, in the specimen in question and Prolibytherium, the mastoid process is in the same
plane with the occipital. In all Giraffidae and presumably in
Progiraffa, these processes are more rostral than the occipital,
which protrudes and forms an hourglass shape. Second,
Prolibytherium and the braincase in question both possess a
deep groove separating the basisphenoid from the temporal
bone, extending from the level of the alisphenoid canal to
the posterior basioccipital tuberosities. This groove is absent
in Cervidae, Bovidae, and Giraffidae. Third, the impression of

The familial affiliation of this Miocene ruminant is currently

unresolved. It is unlikely, however, that Prolibytherium belongs to the Giraffidae (Sivatheriinae or Sivatheriidae;
Hamilton 1973). There are general similarities in the shape of
its cranial appendages to the ossicones of Sivatherium; both
taxa possess massive plate-like cranial appendages that are
fused to the parietal bone. The cranial appendages of
Prolibytherium, however, are not likely to be ossicones, because no sutures are visible separating the cranial appendages
from the skull. In addition, considering differences in the dentition and geologic age, these similarities are more likely due to
convergence. The dentition of Prolibytherium is different from
that of Giraffidae in that the teeth are slender and slightly more
hypsodont, and the enamel is not as crenulated (Hamilton
1973). Sivatherium (~2 Ma) is also significantly younger than
Prolibytherium (~1916 Ma) (McKenna and Bell 1997).
Prolibytherium has been previously assigned to
Palaeomerycidae based on the cranial appendages (Janis and
Scott 1987; Prothero and Liter 2007; Solounias 2007).
Palaeomerycids were geographically widespread in North
America and Eurasia, and had simple supraorbital appendages
with a median single occipital appendage (Ginsburg and Heintz
1966; Qiu et al. 1985; Prothero and Liter 2007). None of the
palaeomerycids have a forward projection-extension of the cranial appendages, as is seen in Prolibytherium. In addition, the
cranial appendages of palaeomerycids have been classified as
frontal ossicones, due to the presence of visible suture lines with
the skull (Astibia et al. 1998; Snchez et al. 2010). The
Prolibytherium cranial appendages, however, are fused to the
parietal bone with no visible suture. The internal structure of
true giraffid ossicones is composed of cartilaginous material as
well as dense connective tissue (Spinage 1968; Ganey et al.
1990). The Prolibytherium cranial appendages, however, appear to consist of cancellous bone surrounded by a thick bony
cortex (Fig. 1b), further negating their identification as
ossicones. Lastly, the Prolibytherium dentition lacks the
BPalaeomeryx fold,^ shared by many Palaeomerycidae
(Prothero and Liter 2007).
Current observations suggest a closer relationship to
Climacoceratidae. Morales et al. (2008) described specializations of the cubonavicular that are shared only between
Climacoceratidae and Prolibytherium. In addition, the
Prolibytherium magnieri cranial appendages are oriented parallel to the dorsal aspect of the skull. This orientation matches that
of the anterior beam of the cranial appendage of Climacoceras
gentryi (Hamilton 1978: Fig. 21). The curvature between the

J Mammal Evol

anterior and posterior beams of Climacoceras gentryi also

matches the dish shape of the Prolibytherium plate-like appendages. The cranial appendages of Climacoceras are similar to
cervids in shape but have no basal burr, negating a cervid type
detachment. Pickford et al. (2001), Morales et al. (2003), and
Snchez et al. (2010) placed Prolibytherium near Climacoceras.
If the presumed cervid-like-shaped cranial appendage from
Gebel Zelten (specimen NHM M 26690) is the female of
Prolibytherium, as Snchez et al. (2010) have proposed, then
the affinity of Prolibytherium to Climacoceratidae is strengthened. However, evidence strongly suggests that early Miocene
female giraffoids would have been hornless (Solounias 2007).
We feel that the specimen described as a female Prolibytherium
(NHM M 26690) is more likely a male representing a new
taxon. We agree, however, with the similarities proposed between the Climacoceras and Prolibytherium cranial appendages.

Functional Implications of the Fused Condyles

The atypical morphology of the occipital condyles in
P. magnieri and P. fusus has functional implications. We hypothesize that the fusion and increase in thickness on the
ventral margin of the occipital condyles allows for more
centralized range of motion and support of the head. The
convergence of the condyles is so complete that there is no
visible fusion line, indicating that the merging happens early
in ontogeny. This thickened increase in central bony material
suggests a median plane focus of forces during combat. The
fusion also provides a greater articular surface area between
the occipital condyles and the atlas during flexion of the head.
Hamilton (1973) also described large anterior swellings on the
basioccipital region of P. magnieri, which according to Mead
(1906), provided strength and support to the atlanto-occipital
joint during combat. In addition, the deep groove extending
from the level of the alisphenoid canal to the posterior
basioccipital tuberosities, which is absent in other ruminants,
appears to isolate the basioccipital bone from the temporal
bone laterally. Presumably, this is congruent with a more centralized modality of fighting. Lastly, the thickened dorsal and
ventral arches of the atlas would provide further support to the
strengthened atlanto-occipital joint.
Presumably, midline ventral fusion of the occipital condyles
would increase the range of motion for rotation of the head.
Conversely, dissections of the atlanto-occipital bone of several
sheep specimens revealed that rotation is limited by a V-shaped
notch on the anterior articular facets of the atlas, which articulates with the junction between the posterior and ventral articular occipital condylar surfaces. This barrier to rotation prevents
the articular surface of the atlas from reaching the midline of the
occipital condyles, regardless of midline fusion. The
Prolibytherium occipital condyles, however, have a rounded
junction between the posterior and ventral articular surfaces,

and a widened notch on the atlas, suggesting that their rotation

range of motion was in fact likely increased.
Relationship of Cranial Appendages to the Specialized
Atlanto-Occipital Joint
The approximation of the occipital condyles provides greater
central support of the head, which is especially important for
species with large and heavy cranial appendages.
Prolibytherium magnieri possesses massive Bbutterfly wing^
shaped appendages encompassing the dorsal surface of the
frontal, parietal, and occipital aspects of the skull (Snchez
et al. 2010). These large cranial appendages are oriented
somewhat parallel to the calvaria, and the surface extends
anterior to the orbit, lateral to the margins of the skull, and
significantly posterior to the occipital. This plate-like orientation of the cranial appendages is a unique feature of
Prolibytherium and is not seen in any extant or extinct ruminant. The exceptionally large and flattened cranial appendages
were likely utilized in a specialized mode of combat requiring
strong reinforcement of the atlanto-occipital joint.
While the ventral fusion of the occipital condyles is a feature unique to Prolibytherium, two specimens of
Schansitherium tafeli (AMNH 32505, HPM 6217) and one
specimen of Giraffokeryx punjabiensis (AMNH 19475) (belonging to Giraffidae) exhibit approximated condyles, with a
narrow intercondylar groove. Both of these giraffids possess
two pairs of ossicones, potentially necessitating stronger support of the atlanto-occipital joint. The Giraffokeryx specimen
has a pair of ossicones on the anterior frontal bone, and a
larger pair of ossicones above the orbit. The posterior
ossicones sit on an expanded boss and extend posterolaterally from the frontal bone (Solounias 2007). Both
Schansitherium specimens have a small pair of ossicones extending in front of the orbit, oriented anteriorly, and large,
straight posterior ossicones situated above the orbit. The occipital condyles of these three specimens are approximated on
the rostral ventral margin, separated only by a narrow groove,
which is especially shallow in one Schansitherium specimen
(HPM 6217) (Solounias 2007). The massive appendages of
Prolibytherium, and the multiple pairs of ossicones seen in
Giraffokeryx and Schansitherium likely require enhanced
cranio-cervical support, contributing to the approximation of
the occipital condyles.
Acknowledgments We acknowledge the NYIT-COM Academic
Scholars program. We thank Pip Brewer and Jerry Hooker at the NHM.
We thank Eileen Westwig, and the Departments of Mammalogy and
Paleontology of the AMNH for access to specimens. We also thank Tao
Deng, Zhanxiang Qiu, Banyue Wang, and Sukuan Hou at IVPP. We thank
John Barry for discussions on the systematics and geologic information.
We also thank Larry Flynn and Michelle Morgan and the Peabody Museum of Harvard. We also thank Everett Lindsay. Funds were covered by
NS.

J Mammal Evol

References
Antoine P-O, Mtais G, Orliac M, Crochet J-Y, Flynn L, Marivaux L,
Rajpar AR, Roohi G Welcomme J-L (2013) Mammalian Neogene
biostratigraphy of the Sulaiman Province, Pakistan. In: Wang X,
Flynn L, Fortelius M (eds) Fossil Mammals of Asia: Neogene
Biostratigraphy and Chronology. Columbia University Press, New
York, pp. 400422
Arambourg C (1961) Prolibytherium magnieri, un vellricorne nouveau
du Burdigalien de Libye. Compt Rendu som Soc Gol France 3:61
62
Astibia H, Morales J, Moy-Sol S (1998) Tauromeryx, a new genus of
Palaeomerycidae (Artiodactyla, Mammalia) from the Miocene of
Tarazona de Aragn (Ebro Basin, Aragn, Spain). Bull Soc Gol
France 169:471477
Barry JC, Cote C, MacLatchy L, Lindsay EH, Kityo R, Rajpar AR (2005)
Oligocene and early Miocene ruminants (Mammalia, Artiodactyla)
from Pakistan and Uganda. Palaeontol Electron 8:122
Ganey T, Ogen J, Olsen J (1990) Development of the giraffe horn and its
blood supply. Anat Rec 227:497507
Ginsburg L, Heintz E (1966) Sure les affinitis du genre Palaeomeryx
(ruminant du Miocne europen). Compt Rendu San Acad Scien
262:979982
Hamilton WR (1973) The lower Miocene ruminants of Gebel Zelten,
Libya. Bull Brit Mus Nat Hist (Geol) 21:76150
Hamilton WR (1978) Fossil giraffes from the Miocene of Africa and a
revision of the phylogeny of Giraffoidea. Phil Trans R Soc (B) 283:
165229
Janis CM, Scott KM (1987) The interrelationships of higher ruminant
families with special emphasis on the members of Cervoidea. Am
Mus Novitates 2893:185
Lindsay EH, Flynn LJ, Cheema IU, Barry JC, Downing KF, Rajpar AR,
Raza SM (2005) Will Downs and the Zinda Pir Dome. Palaeontol
Electron 8:118

McKenna MC, Bell SK (1997) Classification of Mammals above the

Species Level. Columbia University Press, New York
Mead CS (1906) Adaptive modifications of the occipital condyles in
Mammalia. Am Nat 40:475483
Morales J, Soria D, Nieto DM, Pelez-Campomanes P, Pickford
M (2003) New data regarding Orangemeryx hendeyi from the
type locality, Arrisdrift, Namibia. Mem Geol Surv Namib 19:
305344
Morales J, Soria D, Pickford M (2008) Pecoran ruminants from the early
Miocene of the Sperrgebiet, Namibia. Mem Geol Surv Namib 20:
397464
Pickford M, Attia YS, Abd el Ghany MS (2001) Discovery of
Prolibytherium magnieri Arambourg, 1961 (Artiodactyla,
Climacoceratidae) in Egypt. Geodiversitas 23:647652
Pilgrim GE (1908) The Tertiary and Post-Tertiary fresh water deposits of
Baluchistan and Sind with notices of new vertebrates. Rec Geol
Surv India 2:139166
Pilgrim GE (1911) The fossil Giraffidae of India. Palaeontol Indica 4:1
29
Prothero DR, Liter MR (2007) Family Palaeomerycidae. In: Prothero DR,
Foss SE (eds.) The Evolution of Artiodactyls. The Johns Hopkins
University Press, Baltimore, pp 241248
Qiu Z, Yan D, Jia H, Sun B (1985) Preliminary observations on the newly
found skeletons of Palaeomeryx from Shanwang, Shandong.
Vertebr Palasiatic 23:173195
Snchez I, Quiralte V, Morales J, Azanza B, and Pickford M (2010)
Sexual dimorphism of the frontal appendages of the early Miocene
African pecoran Prolibytherium Arambourg, 1961 (Mammalia,
Ruminantia). J Vertebr Paleontol 30:13061310
Solounias N (2007) Family Giraffidae In: Prothero DR, Foss SE (eds.)
The Evolution of Artiodactyls. The Johns Hopkins University Press,
Baltimore, pp 257277
Spinage CA (1968) Horns and other bony structures of the skull of the
giraffe, and their functional significance. E Afr Wildl J 6:5361